Family Tetillidae Sollas, 1886

Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest © Kluwer Academic/Plenum Publishers, New York, 2002

Rob W.M. Van Soest1 & Klaus Rutzler2

'Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT, Amsterdam, Netherlands, ([email protected]) ^ National Museum of Natural History, Smithsonian Institution, Washington DC. 20560, U.S.A. ([email protected])

Tetillidae Sollas (Demospongiae, Spirophorida) are for the most part globular sponges ('golf ball' sponges) belonging to a small order on account of their possession of sigmaspire microscleres and triaene megascleres. The characteristic megasclere is the protriaene, which is found in most members of the family with very few exceptions. In addition there are long oxeas and frequently anatriaenes, rarely also calthrops or amphitriaenes. The skeletal architecture is strictly radial. Eight valid genera are recognized, differentiated by presence of cortical structures, specialized pore-sieves, and composition of the spicule complement. Tetillids are common and ubiquitous sponges in all oceans and at all depths. Keywords: Porifera; Demospongiae; Spirophorida; Tetillidae; Acanthotetilla; Amphitethya; Cinachyra; Cinachyrella; Craniella; Fangophilina; Paratetilla; Tetilla.

DEFINITION, DIAGNOSIS, SCOPE megascleres). The family comprises more than one hundred species distributed over all oceans and habitats. Synonymy Biology [Lophurellidae] Gray, 1872a. [Casuladae] Gray, 1872a. [Tethyina] Carter, 1875b (not Tethyadae Gray, 1848). Tetillidae Tetillids have a preference for sedimented habitats and some Sollas, 1886a. Tethyopsillidae Lendenfeld, 1903. Tethydae species possess a root of long spicule bundles to attach them to the Lendenfeld, 1907. Ectyonillidae Ferrer-Hernandez, 1914a. substrate. Reproductive patterns range from the extrusion of fertilized Craniellidae de Laubenfels, 1936a. eggs, which fix to the substrate and develop directly, to incubation of complete young sponges which are then expelled by localised Definition breakdown of the pinacoderm. No free larvae have yet been described. Spirophorida with sigmaspire microscleres and a radiate skeleton of triaenes and oxeas. Remarks

Diagnosis Gray (1867a) included sponges of this group under the genus name Tethya in a larger Tethyadae comprising a wide range of Typically with spherical growth form, often with characteristic tetractinomorph genera. Carter (1875b) was the first to distinguish pits called porocalices containing the inhalant and occasionally also a separate group (subfamily) for tetillids, but like Gray called the exhalant orifices. Cortical region often strengthened by collagen group Tethyina. The explanation for this is that at that time there fibers and special cortical megascleres, but this region may be thin was still a great confusion over the identity of Tethya. Lamarck or absent in several genera. Skeleton with tetraxonic and monaxonic (1815) included in his genus Tethya next to T lyncurium megascleres (triaenes, huge oxeas) organized in a radiate pattern of ( = T aurantium) and others, Tethya cranium (Miiller, 1789 as spicule bundles, often spiralling outwards from the centre of the Alcyonium). Carter was aware of the great difference between the body; oxeas, protriaenes and anatriaenes are most common and type of Tethya (i.e., T lyncurium) and T cranium (see for example these often protrude from the surface producing a conulose or hairy Carter, 1872a), but he ignored Schmidt's (1870) proposal to surface. In sponges provided with porocalices they form a palisade employ a separate genus Craniella for Alcyonium cranium. of long spicules, protruding far beyond the sponge surface, sur- Schmidt (1870) included Craniella in his family Ancorinidae along rounding the porefields or oscular apertures, which lie in rounded with Stelletta and other astrophorid sponges, but Sollas (1886a) depressions. Microscleres are unique contorted microspined sig- revived Carter's Tethyina under a different name by erecting the maspires. Reproduction is oviparous without a larval stage, or family Tetillidae based on Schmidt's (1868) genus Tetilla. In many viviparous with production of young adults within a parent. ways this was an unwise choice of type genus, inspired by some notion that Tetilla would represent the most primitive of tetillid Scope genera. The description of Tetilla euplocamos by Schmidt (1868: 40, pi. V fig. 10) is insufficient to conclude with certainty what its Out of twenty six nominal genera eight are considered valid, properties were. Selenka (1880: 469) thought to recognize Acanthotetilla, Amphitethya, Cinachyra, Cinachyrella, Craniella, Schmidt's species in an intertidal habitat in the Bay of Rio de Fangophilina, Paratetilla, Tetilla. They are differentiated on gross Janeiro, and provided some additional data. Sollas (1888) was able morphological structure (presence or absence of a cortical region, to verify from a Schmidt slide that at least protriaenes were pres- presence or absence of specialized pore areas), and spicule comple- ent, but apparently this species has no sigmaspires (cf. below). ment (possession of calthrops, amphitriaenes or other auxiliary Even though protriaenes are of common occurrence in tetillids,

85 86 Porifera • Demospongiae • Spirophorida • Tetillidae they are by no means confined to that group and we have to rely and Tethyopsillidae was incorrect, since the type of Tetillidae and on the subtle tetilliform protriaene-type as proof that Tetilla genus Tetilla (T euplocamos) has no sigmaspires and thus would euplocamos is a member of what is generally understood as Tetilla belong in Tethyopsillidae, making this family an objective synonym and the Tetillidae. In summary: we have a type species of the type of Tetillidae. De Laubenfels (1936a) for the same reason refused to genus of a family lacking the major synapomorphy of the order to accept Tetilla euplocamos as a close relative of Craniella, and which the family belongs! Some confidence may be obtained from erected a separate family Craniellidae for the genera possessing sig- the fact that Tetilla species lacking sigmaspires are not uncommon. maspires. Tetilla and Tetillidae were defined as 'Choristida lacking Despite this dubious type species designation, Sollas' (1888) mono- microscleres' and an unrelated group of genera was assigned to it. graphic treatment of Tetillidae was followed by most subsequent Needless to say that neither scheme gained general support. authors with the exception of Lendenfeld (1888, 1903, 1907) and de Laubenfels (1936a). Lendenfeld continued to employ Tethya for Previous reviews sponges answering to the definition of Craniella, but at least kept this in the family Tetillidae. He erected a family Tethyopsillidae for Sollas (1888), Lendenfeld (1903), Wilson (1925), Topsent sponges without sigmaspires. Lendenfeld's division into Tetillidae (1928c), Levi (1973), Rutzler (1987).

KEY TO GENERA

(1) Specialized pore-bearing pits ('porocalices') present 2 No porocalices 6 (2) Spicules include medium-sized heavily spined oxeas ('megacanthoxeas') Acanthotetilla No megacanthoxeas 3 (3) Among the megascleres there are short-shafted triaenes with equal-sized rays (calthrops). N.B. these may have bifid cladi but are not to be confused with long-shafted amphiclads Paratetilla No calthrops, all triaenes - if present - are long-shafted 4 (4) In cross section, there is a clear cortical region with special cortical megascleres (short oxeas) and collagenous reinforcement Cinachyra In cross section there is no discernible peripheral layer different from that of the interior 5 (5) Porocalices all similar, undifferentiated morphologically Cinachyrella Porocalices differentiated into a morphologically distinct inhalant and exhalant type Fangophilina (6) Stalked sponges with differently structured stalk and main body; among the megascleres of the stalk there are triaenes with cladi on both ends (amphitriaenes, -diaenes, -monaenes, collectively called amphiclads) Amphitethya No amphiclads, if stalked, then it is a thin root-like projection, not a proper stiff stalk 7 (7) In cross section, there is a clear cortical region with collageneous reinforcement Craniella In cross section there is no discernible peripheral layer different from that of the interior Tetilla

ACANTHOTETILLA BURTON, 1959 Previous review

Synonymy Van Soest (1977b).

Acanthotetilla Burton, 1959a: 201. Acanthocinachyra Levi, Description of type species 1964b: 386. Acanthotetilla hemisphaerica Burton, 1959a (Fig. 1A-B) Type species Synonymy. Acanthotetilla hemisphaerica Burton, 1959a: 201, fig. 5; Van Soest, 1977b: 2, pi. I figs a-b, pi. II figs c-d, text-fig. 1. Acanthotetilla hemisphaerica Burton, 1959a: 201 (by Material examined. Holotype: BMNH 1936:3:4:530 - John monotypy). Murray Exped. stat. 45, coast of S Arabia, litothamnion bottom, 38 m depth. Definition Description. Semiglobular (Fig. 1 A), 4—5 cm diameter. Colour in alcohol yellowish brown. Surface hispid-bristly due to megascle- Tetillidae with megacanthoxeas as auxiliary megascleres. res projecting 1 mm or more beyond the ectosome. Porocalices numerous, 1-2 mm diameter, scattered over the surface. No appar- Diagnosis ent oscules. Consistency hard, incompressible. Skeleton radiate, forming a tight palisade at the surface. The main megascleres are Globular, hemispherical or irregularly massive sponges. smooth long oxeas and medium-sized megacanthoxeas, but protri- Surface extremely hispid and provided with numerous small-sized aenes and rare anatriaenes are also present. Interiorly, the skeleton porocalices. Skeleton radiating bundles of oxeas, protriaenes, ana- is less dense, making the choanosome cavernous, with megacan- triaenes and megacanthoxeas, developed into an impenetrable pal- thoxeas and sigmaspires as the main spicule categories present. isade at the periphery. Internally cavernous with megacanthoxeas Spicules. Protriaenes, simple, straight, relatively rare, shaft and sigmaspires as the main spicule complement. Four species are 1000-2520 X 6-14 u,m, cladi 30-56 u,m; anatriaenes even rarer, known from the Western Indian Ocean and the Caribbean, depth shaft 1250-1600 X 10 jxm, cladi 55-80 |xm; oxeas, forming the range 38-100 m. bulk of the radiating skeleton, straight, 3000^1400 X 25^10 u,m; Porifera • Demospongiae • Spirophorida • Tetillidae 87

Fig. 1. Acanthotetilla spp. A-B, Acanthotetilla hemisphaerica Rurton, 1959a. A, holotype BMNH 1936.3.4.530 (scale 1cm). B, megacanthoxea of holotype (scale 50 p,m). C, Acanthotetilla enigmatica (Levi, 1964b as Acanthocinachyra), megacanthoxea (scale 50 p,m). D-H, Acanthotetilla gorgonosclera Van Soest, 1977b. D, oxea (scale 10 p,m). E, protriaene (scale lOfjun). F, megacanthoxea (scale 50(jL,m). G, detail of same (scale 10 (xm). H, sigmaspire (scale lfjun). megacanthoxeas (Fig. IB), thickly spined, spines arranged in irreg- 1977b: 7 was described from Barbados (see Fig. 2D-H). It has char- ular 'whorls', with points directed towards the centre, 325^-14 X acters in between A. seychellensis and A. hemisphaerica, and in 40-60 |xm, with 20-25 whorls of spines. Juvenile growth stages of view of its Atlantic occurrence is also considered a valid species. the megacanthoxeas are centrotylote smooth oxeas. Distribution and ecology. Only recorded from the type locality. Remarks. The structure of the skeleton is reminiscent of a AMPHITETHYA LENDENFELD, 1907 cortical specialization (impenetrable palisade of oxeas and megacanthoxeas at the surface), but no organic fibrous cortical region has Synonymy been described so far, so this remains undecided. Burton (1959a) gave a faulty description of A. hemisphaerica by expressly stating Amphitethyahendenfeld, 1907: 126. that sigmaspires were absent. This induced Levi (1964b) to erect a genus Acanthocinachyra for material very similar to Burton's spec- Type species imen, but possessing numerous microscleres. The type species (by monotypy) Acanthocinachyra enigmatica Levi, 1964b: 386, fig. 2, Amphitethya microsigma Lendenfeld, 1907: 126 (by subsequent from Inhaca island, Mozambique (material from MNHN re- designation; Riitzler, 1987, confirmed by Hooper & Wiedenmayer, examined, megacanthoxea figured in Fig. 2C) shares most features 1994: 429). with A. hemisphaerica and since Burton's species (even his own preparations) contain numerous sigmaspires, the two genera are Definition obvious synonyms. The two type species were kept as separate species by Van Soest (1977b), but additional material may bridge Tetillidae with long-shafted amphiclad triaenes and the few discrepancies and it is predicted that they both belong to the plagiotriaenes. same species. A third species from the Western Indian Ocean, Acanthocinachyra seychellensis Thomas, 1973: 80, has unusually Diagnosis thin and curved megacanthoxeas as well as much smaller smooth oxeas, and may indeed be a valid separate species of Acanthotetilla. Globular-stalked sponges with conulose surface, lacking The fourth species, Acanthotetilla gorgonosclera Van Soest, porocalices. Small scattered oscules. Skeleton of the main body 88 Porifera • Demospongiae • Spirophorida • Tetillidae

Fig. 2. Amphitethya microsigma Lendenfeld, 1907. A-D, habit and spicules reproduced from Lendenfeld, pi. XV fig. 19-3. A, habit (scale 1cm). B, spicules. C, detail of amphitriaene. D, detail of sigmaspire (sizes see text). E-H, photos of spicules made from slides of one of the types kept in BMNH. E, cross section of peripheral region of peduncle (scale 500 p,m). F, plagiotriaene (scale 100 p,m). G-H, amphitriaenes (scale 100 u,m).

radiate, consisting of oxeas and normal triaenes, and that of the of Western Australia, 82-110m depth. BMNH 1908.2.9.33-35 stalk longitudinally arranged, consisting of all megascleres includ- (not seen) - 3 slides of one of the syntypes, labelled "Valdivia & ing amphiclad triaenes. Cortical region present in stalk and main Gazelle Lendenfeld". body. Megascleres oxeas, protriaenes, anatriaenes, plagiotriaenes Description (summary of Lendenfeld's 1907 extensive and long-shafted amphiclad triaenes. Microscleres sigmaspires. description). Stipitate sponge (Fig. 2A). Main body globular, up to Two species, both from Australian waters. 4.2 cm in diameter, surface conulose, optically smooth between conules. Stalk up to 7 cm long, 1.2 cm in thickness, circular in cross section, surface also slightly conulose. Colour in alcohol is coffee- Recent review brown. In cross section, both the main body and the stalk show a distinct cortical layer of about 250 u,m thick, in which only Riitzler (1987). microscleres and auxiliary megascleres are found. The amphiclad megascleres are confined to the stalk and are found in the subecto- Description of type species somal layer (Fig. 2E). The main skeleton consists of oxeas, packed longitudinally in the stalk and in radiating bundles in the main Amphitethya microsigma Lendenfeld, 1907 (Fig. 2A-H). body, where they follow a spiral course as is found in many other Synonymy. Amphitethya microsigma Lendenfeld, 1907: tetillids. Megascleres (Fig. 2B), protriaenes, cladome regular, with 126, pi. XV figs 19-39. fusiform shaft, 3000-5000 X 10|xm; anatriaenes, confined to the Material examined. Syntypes: 2MB (not seen) - collected peripheral region of the stalk, 5000-7000 X 10|xm, cladi 15|xm; by the 'Gazelle' Exped., Dirk Hartog Island, off the NW coast plagiotriaenes (Fig. 2F), confined to the peripheral region of the Porifera • Demospongiae • Spirophorida • Tetillidae 89 stalk (Fig. 2E), straight shaft, rounded at the end, 320-1100 X they are mixed with protriaenes. These protrude beyond the 13-30 |xm, cladi 50-250 |xm; amphiclad triaenes (Fig. 2C, G-H), layer of cortical oxeas and are the cause of the bristly surface. confined to the peripheral region of the stalk, shaft 160-540 X Megascleres choanosomal large oxeas, cortical small oxeas, 14-30 |xm, cladome usually irregular with cladi of 100-150 p,m, protriaenes, and anatriaenes. Microscleres sigmaspires. So far, only mostly with different lengths, or occasionally absent (diaene and a single valid species has been recognized with certainty to belong monaene variations are not uncommon); oxeas, straight, or more to this genus, most records of Cinachyra species concern the genus often curved, smooth, gradually but sharply pointed, 6000-8000 X Cinachyrella (cf. below). 20-60 jjLm. Microscleres sigmaspires (Fig. 2B,D), 10-12X0.5-0.8 |xm. Distribution and ecology. Dirk Hartog Island, off the coast of Previous reviews NW Australia, 82-110 m. Remarks. In view of the similarity and the structural position Wilson (1925), Rutzler (1987). they share, the amphiclad triaenes and the plagiotriaenes are very likely to be considered as the same spicule type, in which the Description of type species amphiclad condition may or may not have developed. Lendenfeld assigned to his genus Amphitethya a second tetillid species with Cinachyra barbata Sollas, 1886a (Fig. 3A-C). amphiclad triaenes, Tethya stipitata Carter, 1886c: 460 (redescribed Synonymy. Cinochyra barbata Sollas, 1886a: 183; Cinachyra by Sollas (1888: 49) as Tetilla?). This differs in details of spicule barbata Sollas, 1888: 23, pis III, XXXIX. sizes from A. microsigma, but is otherwise similar and is an Material examined. Syntypes (not seen): BMNH obvious Amphitethya. The fact that both species assigned to 1889.1.1.14-19, 107-108 (slides 1894.11.16.37-51) - the type Amphitethya are prominently stalked sponges is not emphasized series consists of more than 60 syntypes all dredged from the here, as root-organs of other tetillids (e.g., Tetilla euplocamos or shores of Balfour Bay, Kerguelen. Cinachyra barbata) may be considered homologous to a stalk. A Description (from Sollas, 1888). Globular sponges (Fig. 3A), further species assigned by Lendenfeld, viz., Tetilla bacca Selenka, seated on a dense spicular basal mass; ash-grey in alcohol. Oscules 1867, is considered a member of Paratetilla. Its auxiliary spicules inconspicous, scattered. Porocalices large and numerous, up to are calthrops-like, not amphiclad, although the cladi may be occa- 5mm in diameter, scattered over the lateral surfaces, but relatively sionally bifid. Likewise, assignments of Tetilla merguiensis sensu rare on top, and lacking from the basal mass. The porocalices are Topsent, 1897a and Paratetilla aruensis Hentschel, 1912 to flask-shaped, penetrating deep into the interior. Internally, they are Amphitethya by Wilson (1925) are incorrect. The differences are smooth-walled and -rimmed, and have a poresieve at the bottom. perceived such that the Amphitethya amphitriaene is a derivate of a Surface hispid-bristly, with megascleres protruding far beyond the long-shafted triaene (plagiotriaene), whereas the occasional bifid ectosome. Size of main body up to 7X8.5X7 cm, size of basal Paratetilla auxiliary megasclere derives from equal-rayed mass up to 5 X 10 X7cm. Basal mass apparently a fused root sys- calthrops-like spicules. tem which gradually expands during growth of an individual, it is absent in the smallest collected specimen. In cross section (Fig. 3B) the cortical region is visible as a dense peripheral layer, up to CINACHYRA SOLLAS, 1886 1.75 mm in thickness, its whitish colour showing off from the yellowish choanosomal interior. The cortex consists of a distinct Synonymy organic collagenous tissue, reinforced by special cortical oxeas strewn at all angles and microscleres. The choanosomal skeleton [Cinochyra] Sollas, 1886a: 183 (nomen corrigendum); consists of bundles of oxeas issuing from an eccentrically located Cinachyra Sollas, 1888: 23. focus and following a spiral course towards the periphery. The peripheral bundles contain a mixture of oxeas and triaenes, with the Type species latter protruding far beyond the surface. Megascleres (Fig. 3C). Protriaenes, occasionally prodiaenes, in two distinct size categories, Cinochyra barbata Sollas, 1886a: 183 (by monotypy). large structural ones, fusiform, shaft up to 13,000 X 30 |xm, cladi up to 180X16|xm, and small, sinuous, hair-like, up to 130X4u,m, Definition with cladi 16-30 jjim; anatriaenes, occurring only in the basal mass, where they function as anchoring spicules, up to 40,000 X 24 u,m, Tetillidae with cortex reinforced by auxiliary oxeas, with cladi up to 215X28 p,m; oxeas, in two distinct size categories, large flask-shaped porocalices. choanosomal ones, sharply pointed, straight or curved, up to 8000 X 70 jjum; small cortical oxeas, fusiform, bluntly pointed, up to Diagnosis 900X36|xm. Microscleres sigmaspires, 12-16 pjn. Distribution. Kerguelen, Patagonia, Antarctica, sediment-rich bottoms, 45-549 m. Globular sponges with surface covered by numerous porocal- Remarks. The genus name has been widely used by many ices. Full-grown specimens develop an irregular basal mass as authors for sponges now recognized to belong in a separate genus wide as the globular upper part, equivalent perhaps to a stalk. Cinachyrella Wilson, 1925 (cf. below). This shares the porocalices, Porocalices large and deep, flask-shaped, surrounded by a fringe of but lacks any cortical specialization. Sollas (1888) maintains that long megascleres. In cross section there is a prominent cortex some of the porocalices are exhalant, whereas others are inhalant. strengthened by special cortical oxeas. Skeleton of the globular However, Kirkpatrick (1905) found small, simple oscules separate part radiate, consisting of bundles of oxeas originating in a central from the complicated porocalices, and similar findings are reported nucleus and spirally curving outwards toward the periphery, where by Boury-Esnault & Van Beveren (1982) and Rutzler (1987). 90 Porifera • Demospongiae • Spirophorida • Tetillidae

Fig. 3. Cinachyra barbata Sollas, 1888, drawing of habit (A), peripheral skeleton in cross section (B), and spicules (C), reproduced from Sollas' pi. Ill) (scales see text).

CINACHYRELLA WILSON, 1925 Synonymy. Tetilla hirsuta Dendy, 1889: 75; Cinachyra hirsuta; Lendenfeld, 1903: 28; Tetilla (Cinachyrella) hirsuta; Synonymy Wilson, 1925: 363. Material examined. Lectotype (here designated) (not seen): [Psetalia] Gray, 1873c: 234 (nomen oblitum). [Labaria] Gray, BMNH 1889.1.21.22 - (the larger of the two specimens), 1873c: 235 (nomen oblitum). Cinachyrella Wilson, 1925: 363. Rameswaram Island, Gulf of Manaar, Sri Lanka, Thurston collec- Raphidotethya Burton, 1934a: 526. Uliczka de Laubenfels, 1936a: tion. Paralectotype (not seen): BMNH 1925.11.1.769 - the smaller 174. specimen from the same locality. Description (from Dendy, 1889). Globular sponge, size up Type species to 5 cm in diameter, colour dark grey in alcohol. Surface 'hirsute' due to projecting spicules. Porocalices scattered irregularly over Tetilla hirsuta Dendy, 1889: 75 (by subsequent designation; the body. Some porocalices are hemispherical and shallow, others Rutzler, 1987). deep and tubular. The floor of the porocalices is alternatively perforated by a number of small pores, or some oscular tubes, or is Definition devoid of openings. Dendy assumed the porocalices were either inhalant or exhalant, but these are not morphologically differenti- Tetillidae with undifferentiated porocalices, without cortex, ated. In cross section there is no special cortical skeleton. The without auxiliary megascleres. choanosomal skeleton consists of stout radiating fibers issuing from a dense central nucleus. Megascleres. Protriaenes, often Diagnosis of hair-like dimensions, i.e., comparatively thin and short, up to 460 X 14|xm, cladi 50X7 jjum; anatriaenes similar in dimension, Globular sponges with hispid-bristly surface provided with slightly shorter; oxeas, fusiform, sharply pointed, up to 3500 X numerous scattered porocalices; oscules inconspicuous. Skeleton 42 |xm. Microscleres the usual sigmaspires, up to 22 p,m. radiate, consisting of bundles of oxeas issuing from the centre Choanocyte chambers 20|xm in diameter. Distribution. Known and running spirally to the surface. No cortical specialization. only from the type locality. Megascleres protriaenes, anatriaenes, occasionally plagiotriaenes, Remarks. No images of the type species are currently and dominant large oxeas. Microscleres sigmaspires, which may be available, but it is likely that it is closely similar to e.g., Cina- lacking occasionally, finely spined oxeas and/or raphides occur. chyrella apion (Uliczka, 1929 as Cinachyra), as described and More than one hundred species distributed in shallow water of illustrated by Rutzler & Smith (1992). Their illustrations are tropical and subtropical seas. here reproduced in Fig. 4 (habit and surface characteristics) and Fig. 5A (spicules). Recent review Burton (1934a) attempted to revise a large amount of Cinachyrella species (as Cinachyra), arriving at the remarkable con- Rutzler (1987). clusion that most tropical representatives are members of a single intertropical species to be called Cinachyra australiensis (Carter, Description of type species 1886b). Needless to say that such a conclusion is unacceptable. The genus Psetalia Gray, 1873c: 234 was erected for type Cinachyrella hirsuta (Dendy, 1889). species P. globulosa Gray, 1873c: 234 (by monotypy). There are Porifera • Demospongiae • Spirophorida • Tetillidae 91

2 mm

Fig. 4. Cinachyrella apion (Uliczka, 1929), habit and surface characteristics, reproduced from Riitzler & Smith (1992: fig. 6). A, specimen in situ attached to alga, showing numerous buds. B, alcohol-preserved specimen. C, close-up of buds. D, close-up of porocalyx. E, close-up of oscule.

apparently four specimens of this species from Singapore. These are recognizable. The main genus character is the absence of triaenes, globular sponges of up to 6 cm diameter, covered by prominent instead of which there are only oxeas as megascleres, arranged in "tubercles each containing a tuft of filamentous spicules". A deep the usual radiating bundles. At the surface and throughout the concavity is found at the upper surface. It seems clear that this is choanosome there are oxeas of 730 X 12 |xm about half the length a description of porocalyces, and as its type locality is in the tropics, of the choanosomal main oxeas which measure 1400 X 24 |xm. it seems a safe conclusion that it concerns a species of Cinachyrella. Microscleres are sigmaspires apparently in two size categories, The name Psetalia is an unused name in the sense of ICZN Article 24 and 9 \ua, and raphides also in two size categories, 30 and 23.9 (not used after 1899) and accordingly it is proposed to suppress 15|xm. Despite the fact that no porocalices were described, this it in favour of the prevailing junior name Cinachyrella. genus is considered a junior synonym of Cinachyrella, as it has no The genus Labaria Gray, 1873c: 235 was erected for type conulose surface, is hispid, and lacks a cortex. The absence of tri- species Labaria hemisphaerica Gray, 1873c: 235 (by monotypy). aenes and the possession of spicule size categories is not of generic Like Psetalia, this hemispherical sponge of 5 cm diameter appears value, as there is a large diversity in these features among the many to be a Cinachyrella, as Gray described structures on the surface - Cinachyrella species. Moreover, Hooper (pers. comm.) has "... cylindrical perforations, from the centre of which are emitted observed specimens possessing triaenes with their cladomes stick- tufts of elongated filiform spicules..." - which can only be ing some distance beyond the surface. This may also have been the porocalices. The name Labaria is an unused name in the sense of case for the type specimen. ICZN Article 23.9 (not used after 1899) and accordingly it is pro- The genus Uliczka de Laubenfels, 1936a: 174 was erected posed to suppress it in favour of the prevailing junior name for type species Cinachyra schistospiculosa Uliczka, 1929: 45, Cinachyrella. The name is in any case preoccupied by Labaria figs 27-30 from Barbados (by original designation). This species Carter, 1873c (Hexactinellida, Pheronematidae). is a commonplace Cinachyrella, assigned to the synonymy of The genus Raphidotethya Burton, 1934a was erected for type Cinachyrella kuekenthali (Uliczka, 1929 as Cinachyra) by Riitzler & species (by monotypy) Raphidotethya enigmatica Burton, 1934a: Smith, 1992:154. It possesses occasional 'split' oxeas, but these are 526, fig. 2 (here reproduced as Fig. 5B). The holotype (BMNH obvious artefacts, and certainly not a specific or generic character 1930.8.3.27; not examined) is described as a ficiform sponge of (see also Van Soest & Sass, 1981 and Riitzler & Smith, 1992). 4.5 X 2 cm, with a thick stalk and a uniformly hispid surface in Wilson (1925) erected Cinachyrella as a subgenus of Tetilla, which no pores or oscules were visible. No cortical specialization rather than of Cinachyra, thus apparently assuming the lack of 92 Porifera • Demospongiae • Spirophorida • Tetillidae

Fig. 5. A, Cinachyrella apion (Uliczka, 1929), drawing of spicules, reproduced from Riitzler & Smith (1992: fig. 7). B, Cinachyrella enigmatica (Burton, 1934a as Raphidotethya), drawing of cross section and spicules reproduced from Burton's fig. 2 (scales see text). a cortex is a more important feature than the presence of porocalices. CRANIELLA SCHMIDT, 1870 The presence or absence of a cortical specialization is of dubious phylogenetic significance, as it occurs both in tetillids with poro- Synonymy calices (Cinachyra and Cinachyrella) and in those without (Craniella and Tetilla). Moreover, the cortical specialization is also Craniella Schmidt, 1870: 66. Tethya of authors (e.g., Carter, used as a discriminatory character in other tetractinomorph groups. 1872a; Lendenfeld, 1903) (not Tethya Lamarck, 1815). Polyurella Cortical specialization is rather easily detected when the cortex is Gray, 1870b: 312. [Lophurella] Gray, 1872a: 471 (invalid name). thick and strengthened by special cortical megascleres, but remains [Lophiurella] Gray, 1873c: 234 (lapsus). Tethyopsilla Lendenfeld, difficult to demonstrate when it is thin and cortical spicules are 1888: 45. Craniellopsis Topsent, 1913a: 14. lacking. In the case of Cinachyra, the cortex is well-developed, but so far only a single species is known, whereas the bulk of the Type species species described as Cinachyra are devoid of a cortex apparently, and need to be assigned to Cinachyrella. We maintain here the dis- Craniella tethyoides Schmidt, 1870: 66 (by subsequent desig- tinction between the two, because Cinachyra barbata differs also nation; de Laubenfels, 1936a: 175). quite strongly from Cinachyrella hirsuta in spicule sizes and categories. A further complication for the use of Cinachyrella is the Definition relationship with Fangophilina Schmidt (cf. below). This is maintained as a separate genus of Tetillidae on the strength of the occur- Tetillidae without porocalices, with a distinct cortex strength- rence of clearly morphologically differentiated oscular and ostial ened by special cortical oxeas. porocalices. Still, Dendy (1889) examined the bottom of several porocalices in C. hirsuta and distinguished oscular and poral types. Diagnosis Wilson (1925) described special simple oscules from the Philippine specimen he identified as C. hirsuta, and thus casted Globular sponges with conulose but optically smooth surface some doubt even though he acknowledged the possibility that over most of the upper body; at the base there are bundles of Dendy may have been right. It is also not quite certain that Dendy's spicules acting as a root. Oscules few, usually on the top. Ostia in and Wilson's material was conspecific, because there is a large size sieve-like groups overlying subdermal cavities. In cross section discrepancy in the protriaenes recorded from the Indian and there is a distinctly visible cortical layer, which is perhaps divisible Philippine specimens. Later on, Riitzler (1987) expressed strong in a tightly collagenous layer strengthened by radially or confus- doubts about the reality of the specialization of differentiated oscu- edly arranged special cortical megascleres, and a less dense outer lar and ostial porocalices, maintaining that all porocalices are layer in which there are many subdermal cavities. Choanosomal inhalant and that separate simple oscules remain usually unde- skeleton radiating in spiral fashion, with bundles of oxeas originat- tected in most Cinachyra and Cinachyrella. If Riitzler is right and ing from a central focus and spiralling outwards towards the sur- the differentiation will be demonstrated to be an artifact, then face where they are mixed with protriaenes to protrude in groups Fangophilina would be a synonym of Cinachyrella. and push up the ectosome into conical elevations. Megascleres, Porifera • Demospongiae • Spirophorida • Tetillidae 93

Fig. 6. Craniella tethyoides Schmidt, 1870. A, BMNH slide of lectotype. B-D, photos of spicules from lectotype (scales 50p,m). B, sigmaspires. C, protriaene. D, anatriaenes.

protriaenes, anatriaenes, choanosomal oxeas, shorter cortical is sufficient to recognize the genus as valid and of common occur- oxeas. Microscleres, sigmaspires (may be lost not infrequently). rence. The genus is similar in many aspects to Tetilla, but this About 25 species are recorded from most seas and oceans, often genus lacks a cortical specialization. from somewhat deeper localities. The genus Polyurella Gray, 1870b: 312 was erected for type species Polyurella schmidtii Gray, 1870b: 312 (by monotypy). Previous reviews Gray (1870b) admits himself that this is the same (i.e., a junior synonym) as Tetilla (spelled as Tetella (sic)) polyura Schmidt, Sollas, 1888;Topsent, 1894d; Wilson, 1925. 1870: 66, pi. VI fig. 8. This species is here assigned to Craniella on account of the presence of a cortical skeleton of smaller oxeas Description of type species (based a.o. on subsequently assigned specimens in the collection of ZMA). Gray's (1870b) article was published after the appearance Craniella tethyoides Schmidt, 1870 (Fig. 6A-D). of Schmidt's monograph, so Polyurella is a clear junior synonym Synonymy. Craniella tethyoides Schmidt, 1870: 66, of Craniella Schmidt, 1870. pi. VI fig. 9; Sollas, 1888: 54; Tetilla tethyoides; Lendenfeld, The genus [Lophurella] Gray, 1872a: 461 was erected for type 1903: 25. species Tetilla lophura "Schmidt, table" (Gray's notation). There is Material examined. Lectotype (here designated): BMNH no record of Tetilla lophura in any of Schmidt's papers; and it is 1939.2.10.30 (not seen), slide examined BMNH 1870.5.3.54 - also not mentioned in Desqueyroux-Faundez & Stone (1992). We labeled in Schmidt's handwriting "Craniella tethyoides n.g. (116), must assume this is a misspelling, most likely of Tetilla polyura, Florida 100-230 fathoms" (here reproduced in Fig. 6A), presum- since the definition of the family [Lophurellidae] Gray, 1872a: 460 able made from the lectotype which was apparently obtained in an to which this genus is assigned mentions the characteristic numer- exchange with MCZ. The latter museum is presumably holding ous rooting spicules, used by Schmidt to characterize Tetilla a further specimen (here designated paralectotype). Other material. polyura. This is confirmed also by Gray (1873c: 234), where he BMNH 1870.5.3.55 (slide) - Iceland, mentioned in Schmidt's indicates subsequently that Tetilla polyura is the type of his genus report, and not considered part of the type series as it appears to be [Lophiurella] (sic!). Even so, [Lophurella] is not an available genus from a different species. Two further slides from Florida in BMNH name as it cannot be unambiguously assigned to a nominal species- bearing this name on the labels are likewise not part of the type group taxon (ICZN Art. 12.2.5), which rules out type species material. assignment by indication (pers. comm. by Dr Alice Wells of the Description (from Schmidt, 1870, and Sollas, 1888). Australian Biological Resources Survey, Canberra). Likewise, Globular, with conulose surface (where surface membrane is still [Lophurellidae] Gray, 1872a is unavailable as a family name as it is intact). Distinct fibrous cortex strengthened by cortical oxeas based on an unavailable genus name. arranged radially. Choanosomal oxeas robust, forming the bulk of The genus Tethyopsilla Lendenfeld, 1888 was erected (by the megascleres and protruding beyond the surface. Megascleres. monotypy) for type species Tethyopsilla steward Lendenfeld, 1888: Protriaenes (Fig. 6C) present (no size recorded), anatriaenes 45, to accomodate species lacking sigmaspire microscleres. The (Fig. 6D) with cladi of 70-90 |xm, cladome 80-110 pim in diameter type (which was recently found to be missing from the (no length recorded); oxeas in two distinct size categories (no sizes 2MB, and also not located elsewhere, cf. Hooper & Wiedenmayer, recorded). Microscleres (Fig. 6B), sigmaspires up to 35 p.m. 1994: 431) is described as a small (2cm) spherical sponge, with Distribution. Florida, down to 400 m. smooth surface and slit-like oscules. There are abundant protri- Remarks. The type species remains ill-known and awaits aenes, prodiaenes and promonaenes (up to 2000 X 10 pun), anatri- proper redescription from type and fresh material. Nevertheless, the aenes (no size quoted) and oxeas (1500 X 10 pun), no microscleres. combination of the lack of porocalices and a cortical specialization The genus was made the type of a family Tethyopsillidae apparently 94 Porifera • Demospongiae • Spirophorida • Tetillidae originally founded on the possession of irregular protriaenes, but Description of type species later (Lendenfeld, 1903) on the absence of sigmaspires. The poly- mastiid genus Proteleia Dendy & Ridley, 1886 was also included Fangophilina submersa Schmidt, 1880b (Fig. 7A). in this nominal family. The absence or rarity of sigmaspires is Synonymy. Fangophilina submersa Schmidt, 1880b: 73, rather a common phenomenon in unrelated tetillids and not worthy pi. X fig. 3; Topsent, 1920a: 2; Cinachyra submersa; Lendenfeld, of genus or family distinction. Wilson (1925) assigned Tethyopsilla 1903: 28. to Tetilla, but Lendenfeld (1903) expressly mentions a cortex of Material examined. Lectotype (here designated) (not seen): 1.2 mm thickness (admittedly without special cortical oxeas), so it ZMUS P0160 - dry, Caribbean. Paralectotypes: ZMUS P.0166 - seems more appropriate to assign the genus to Craniella. Hooper & dry fragments. Fragments of types: 2MB 6650 (6 slides) - fide Wiedenmayer (1994) remarkably assigned T. stewarti to Cinachyra, Desqueyroux-Faundez & Stone, 1992. but this seems unlikely in view of the smooth surface and slit-like Description (from Schmidt, 1880b and Topsent, 1920a). oscules. The type redescribed by Topsent represents half of the original The genus Craniellopsis Topsent, 1913a was likewise erected specimen. It is 3.4cm high and has a diameter of 3.8 cm. It has no for tetillids possessing a cortex and lacking sigmaspires. Topsent trace left of the spicular root mentioned in Schmidt's description. (1913a) named three species, of which de Laubenfels (1936a: 171) The surface is hispid and many foreign objects such as subsequently designated Tetilla zetlandica (Carter, 1872a: 417) as foraminifera and sand grains have attached to it. At opposite sides the type species. Four syntypes (BMNH 1983.6.14.1^4), as well as there are two large porocalices, both with palisade of very long several slides made from these (fide Hooper & Wiedenmayer, spicules protruding up to 1.5 cm above the surrounding sponge sur- 1994: 432), are extant but have not been examined for this study. face. The porocalices are 1-1.5 cm deep and 0.8-1 cm in diameter. This is a common species in the North Atlantic, sharing the two The left one has a perforated bottomplate, the other shows a few size categories of oxeas with C. tethyoides, the smaller category of slit-like apertures in the bottom and the sides, but is generally with- which is concentrated in the cortex. The lack of sigmaspires is con- out visible apertures. Schmidt apparently misinterpreted the func- sidered of low importance. Lendenfeld (1903) and de Laubenfels tions of both types, but this was corrected by Topsent: the one (1936a) maintained a separation at the genus and family level depicted left in Schmidt's drawing (here reproduced as Fig. 7A) for species with (Craniellidae de Laubenfels) and without sig- is the inhalant porocalyx, the other the exhalant porocalyx. The maspires (Tetillidae sensu de Laubenfels, Tethyopsillidae spicule palisade is made up of oxeas and protriaenes/prodiaenes. In Lendenfeld) (cf. above). cross section the structure of the sponge - apart from the porocalices - is radiate, with bundles of oxeas originating from a focal point traversing towards the surface. There is no cortical specialization. Megascleres. Protriaenes and prodiaene modifications, FANGOPHILINA SCHMIDT, 1880 length not recorded, thickness of shaft up to 20 |xm, cladi up to 400 |xm long, cladome 130|xm wide; anatriaenes, length not recorded, Synonymy thickness up to 8jjim, cladi up to 47jjim, cladome 27jjim; plagio/ orthodiaenes and -monaenes, length not recorded, thickness 25 |xm, Fangophilina Schmidt, 1880b: 73. Spongocardium Kirkpatrick, cladi, sometimes flexuous, up to 420 u,m; oxeas in two size cate- 1902: 224. gories, largest 25000 X 75 |xm, smallest 900-1500 X 15-30 ^m (the latter are found in the choanosome intercrossing the bundles of Type species oxeas). Microscleres sigmaspires 15-35 p,m, larger ones often elongated, wavy, toxiform. Fangophilina submersa Schmidt, 1880b: 73 (by subsequent Remarks. Although he did not define it, Schmidt (1880b) designation; Rutzler, 1987: 189). apparently considered Fangophilina to comprise a wider group of sponges than only Fangophilina submersa, since he assigned Definition Tetilla polyura Schmidt, 1870, Tetilla euplocamos Schmidt, 1868, and Tetilla radiata Selenka, 1880 to it in the discussion of the new Tetillidae with morphologically distinct exhalant and inhalant genus. He apparently forgot that Tetilla had been erected earlier by porocalices positioned laterally, without cortical specialization, himself with Tetilla euplocamos as the type. Thus, we may surmise without auxiliary megascleres. that Fangophilina was intended for at least several tetillids that did not share the peculiar porocalyx-specialization of F. submersa. The Diagnosis genus is of rather dubious validity, since several authors (e.g., Rutzler, 1987) have expressed doubts about the nature of the exha- Globular sponges with hispid surface provided with two large lant porocalyx. Possibly, true oscules have become obscured porocalices situated laterally. These are differentiated into an exha- through contraction in preserved state. Still, the fact that several lant and an inhalant porocalyx. Skeleton radiate, with strongly pro- species have been described with similar structure is considered truding megascleres. Cortical specialization absent. Megascleres sufficient to maintain the genus as valid for the time being (there is protriaenes, anatriaenes, oxeas. Microscleres sigmaspires. Three also the consideration, that Fangophilina is senior to Cinachyrella, species have been assigned to this genus, distributed in the the genus most similar to it). Caribbean and Indian Ocean. Kirkpatrick, 1902, erected Spongocardium for type species (by monotypy) Spongocardium gilchristi Kirkpatrick, 1902: 224, Previous reviews pi. II fig. 1, pi. Ill fig. 1. This sponge (Fig. 7B) is very similar to Schmidt's Fangophilina submersa, with the poral and oscular Lendenfeld (1907); Topsent (1920a); Wilson (1925). porocalyx in a lateral position. There are a few spicular differences Porifera • Demospongiae • Spirophorida • Tetillidae 95

Fig. 7. Fangophilina species. A, Fangophilina submersa Schmidt, 1880b, drawing of cross section of holotype reproduced from Schmidt's pi. X fig. 3 (scale 1 cm). B-C, Fangophilina gilchristi (Kirkpatrick, 1902 as Spongocardium). B, lectotype and cross section, reproduced from Kirkpatrick's pi. II fig. 1 (scale 1 cm). C, drawing of spicules from Kirkpatrick's pi. Ill fig. 1 (scale see text).

(Fig. 7C) (e.g., there are orthotriaenes, 4900 X 35 |xm and a second Diagnosis smaller category of protriaenes, 5300 X 31 and 690 X 2 p,m). Other measurements are similar, oxeas up to 10mm X 80-100|xm, anatri- Globular sponges with hairy-bristly surface, provided with aenes ditto, sigmaspires 35-45 pjn. However, Topsent's conclusion porocalices. Skeleton radiate, with bundles of oxeas originating that Spongocardium gilchristi is a junior synonym of Fangophilina from a central focus and running radially to the surface. Spicules submersa appears incorrect in view of the considerable differences. protriaenes, anatriaenes, calthrops-like triaenes, oxeas, sig- Later, Kirkpatrick (1905) realized that Schmidt's genus was maspires. So far only a single wide-spread Indo-West Pacific defined on the same characters, and pronounced the two genera species is reliably recorded. synonyms. This was followed by Lendenfeld (1907: 157), who erected a third species of Fangophilina, F. hirsuta. Previous reviews

Rutzler(1987).

PARATETILLA DENDY, 1905 Description of type species

Synonymy Paratetilla bacca (Selenka, 1867) (Fig. 8A-C). Synonymy. Stelletta bacca Selenka, 1867: 569, figs 14-15; Paratetilla Dendy, 1905: 97. Paratetilla bacca; Hooper & Wiedenmayer, 1994: 433, with additional synonyms. Tethya merguiensis Carter, 1883a: 366, pi. XV Type species figs 6-8; Tetilla merguiensis; Sollas, 1888: 14. Tetilla ternatensis Kieschnick, 1896: 527. Paratetilla cineriformis Dendy, 1905: 97, Tethya merguiensis Carter, 1883a: 366 (by subsequent desig- pi. Ill fig. 7. nation; de Laubenfels, 1936a: 175). This is generally considered a Material examined. Type material: not located - Samoa. Type junior synonym of Stelletta bacca Selenka, 1867. material of T. merguiensis. IM (presumed) - Burma (=Myanmar). Other material. ZMA specimens - Indonesia, 'Siboga' and 'Snellius Definition IF collections (subjectively assigned to this species). Description (based on Carter, 1883a and Selenka, 1867). Tetillidae with porocalices, with megascleres including Globular (Fig. 8B), size 2-3 cm diameter (larger in subsequently calthrops-like short-shafted triaenes. recorded specimens), surface uniformly hispid due to projecting 96 Porifera • Demospongiae • Spirophorida • Tetillidae

Fig. 8. Paratetilla bacca (Selenka, 1880) (scales see text). A, drawing of spicules reproduced from Selenka, 1867: fig. 15, as Stelletta). B-C, drawing of habit (B) and spicules (C) of Tetilla merguiensis Carter, 1883a: pi. XV figs 6-8, considered a junior synonym.

megascleres, with numerous larger apertures, 0.5-1 cm diameter, Cinachyrella. The calthrop-like short-shafted triaenes are probably scattered evenly over the surface in Selenka's specimen, concen- derived from plagiotriaenes, which are occasionally recorded in trated in the median region in Carter's specimen. Both authors Cinachyrella (cf. Riitzler, 1987; Riitzler & Smith, 1993), and thus interpret these as exhalant, but it is obvious they are porocalices. could be interpreted as a species rather than a genus character. Colour: black-brown (according to Carter). In cross section there is no cortex (Selenka mentions a thin cortical layer containing a layer of calthrops, but this is simply the peripheral region), and the skele- TETILLA SCHMIDT, 1868 ton is radiate, with bundles of oxeas issuing from a central focus. Megascleres (Fig. 8A, C; sizes combined from data and drawings Synonymy of Selenka, Carter, Sollas and Dendy). Protriaenes, up to 3500 X 15 (Jim, cladi up to 90u,m; anatriaenes 3500 X 15p,m, cladi 60|xm; [Dactylella] Gray, 1872a: 461 (nomen oblitum). [Casula] calthrop-like short-shafted triaenes (Selenka: 'vierzackige Sterne'; Gray, 1872a: 461 (nomen oblitum). Tetilla Schmidt, 1868: 40. Carter: 'zone-spicule', Sollas: 'orthotriaenes'), cladi 100-200 u,m Chrotella Sollas, 1886a: 181. Spiretta Lendenfeld, 1888: 42. (Carter gives l/56th of an inch, which would be 450 |xm, clearly Trachygellius Topsent, 1894c: 8. Ectyonilla Ferrer-Hernandez, in excess of the sizes found in other recorded specimens); oxeas, 1914a: 452. Kaira de Laubenfels, 1936a: 175. up to 7000 X 42 |xm. A smaller category of oxeas is mentioned by Carter (1883a) and Dendy (1905 as Paratetilla cineriformis). Type species Microscleres sigmaspires (not recorded by Selenka), ll-17|xm. Distribution and ecology. Widespread Indo-West Pacific (Samoa, Tetilla euplocamos Schmidt, 1868: 40 (by monotypy). Myanmar, Indonesia, NE Australia, Sri Lanka), reefs, and deeper water, down to +200 m. Definition Remarks. Riitzler (1987) nominated Paratetilla cineriformis Dendy, 1905 as the type species, but this was superseded by Tetillidae without porocalices, without cortical specialization, de Laubenfels' (1936a) previous nomination of Tethya merguiensis without auxiliary megascleres. as the type. Since this species was mentioned by Dendy as a species of Paratetilla, de Laubenfels' designation is valid. All Diagnosis described tetillid species with calthrop-like short-shafted triaenes appear to be referable to a single widespread species, Paratetilla Globular sponges with conulose or uniformly hispid surface, bacca (cf. synonymy in Hooper & Wiedenmayer, 1994). If this is with few oscules, usually situated at the top. Usually with spicule confirmed in comparative studies, the validity of this genus is dubi- strands at the base, acting as root system. Surface without porocal- ous, because the possession of the genus character is then limited ices. In cross section, there is no visible cortex. The skeleton is spi- to a single species, which in all other aspects conforms to rally radiate, with bundles of oxeas originating from a central focus Porifera • Demospongiae • Spirophorida • Tetillidae 97

Fig. 9. Tetilla euplocamos Schmidt, 1868. A, drawing of type and protriaene reproduced from Schmidt, 1868: pi. V fig. 10 (sizes see text). B, BMNH slide of type. C-D, photos of spicules from BMNH slide. C, oxeas and bundle of small? protriaenes (scale 100 (xm). D, protriaene (arrow) (scale 100 p,m). and radiating in a spiral manner to the surface. At the periphery, (Fig. 9C, D) in photos made from the BMNH slide. Distribution these bundles of oxeas become mixed with triaenes, predominantly and ecology. Known only from type locality. protriaenes. Microscleres sigmaspires (absent in the type species). Remarks. For no apparent reason, Sollas (1888) changed About 50 species have been described from all parts of the world euplocamos into euplocamus. The species remains ill-known. oceans, mostly from deeper localities. Since many authors did not Schmidt's description is casual and short and subsequent speci- distinguish between Tetilla and Craniella, the actual number of mens from the same area identified under this name (Selenka, species may be less. 1880) are only presumably conspecific. Nevertheless, there are many species of tetillids sharing the characters described by Previous reviews Schmidt, and the genus is considered clear and unequivocal. The genus Dactylella Gray, 1872a: 461 was erected for type Sollas (1888); Rutzler (1987). species Tethya dactyloidea Carter, 1869a: 15, figs \-A from the SE coast of Arabia (by original designation). This is undoubtedly a Description of type species member of Tetillidae, because Carter described a radiate structure and oxeas and protriaenes as megascleres. No certainty exists about Tetilla euplocamos Schmidt, 1868 (Fig. 9A-D) the presence of sigmaspires, because Carter had "given away the Synonymy. Tetilla euplocamos Schmidt, 1868: 40, pi. V specimen" (to Dr Bowerbank, cf Carter, 1871e: 103). In view of fig. 10; Selenka, 1880: 469, pi. 27 fig. 5. the absence of clear porocalyces and apparent lack of a cortical Material examined. Holotype (not seen): MZUS P0206 - skeleton, this is likely to be a Tetilla. Since the name Dactylella Desterro, Bay of Rio de Janeiro, intertidal, coll. F. Mullen Slide of predates Tetilla it needs to be suppressed as an unused name in the the holotype (examined; Fig. 9B): BMNH 1868.3.2.32 - bearing sense of ICZN Article 23.9. The name Tetilla has been frequently Schmidt's handwriting, labeled "Desterro/Tetilla euplocamos Sch used in the past 50 years by many authors, and thus fulfils the 1868". Fragment 2MB 7165 - (see Desqueyroux-Faundez & demands for continued usage under Article 23.9. The name Stone, 1992). Dactylella has not been used for a tetillid sponge after its original Description (from Schmidt, 1868). Pear-shaped (Fig. 9A), proposal, although subsequently, [Dactylella] was also used by with long root ('Nadelschopf'). No cortex. Surface finely hispid. Thiele, 1898 for type species Dactylella hilgendorfi Thiele, 1898: In cross section the choanosomal skeleton consists of bundles of 56, pi. 4 fig. 8, pi. 5 fig. 25, pi. 8 fig. 41a-b. This is a sponge of the oxeas originating from a core spiralling outwards toward the sur- family Dictyonellidae, and the name is now considered preoccu- face, where they mix with triaenes. The latter protrude beyond the pied, and replaced by Lipastrotethya de Laubenfels, 1954. surface and cause a fine downy pelt of spicules. Megascleres. The genus Casula Gray, 1872a: 461 was erected, in Casuladae Protriaenes (Fig. 9A, no size recorded), apparently no anatriaenes; Gray, 1872a: 461, for Tethya casula Carter, 1871e: 99, pi. IV from oxeas 2300 X 22 |xm. No sigmaspires were detected in the slide, nor South Africa (by monotypy). The description of Carter leaves no recorded by Schmidt or others. Some spicules are here illustrated doubt that this is a Tetilla, probably considerably damaged during 98 Porifera • Demospongiae • Spirophorida • Tetillidae collection giving rise to the rather strange schematic drawing of the The genus Trachygellius Topsent, 1894c: 8 was erected for habit in his fig. 1. In its turn this induced Gray (1872a) to empha- type species Trachya globosa Carter, 1886a: 121 from shallow size in his definition of the family Casuladae, that the sponge is water off Port Phillip Head, Victoria (by original designation). This free-living and has a "funnel-shaped expansion or disc formed of is a yellow globular sponge with a short stalk and uneven surface. elongated spicules united together". Spiculation include protri- Oscules in a group around the top. The species is described by aenes up to 10 mm, oxeas up to 5 mm, and sigmaspires of 14 p.m. Carter (1886a) as having a skeleton that is radiate with a "thick der- Since the family group name Casuladae Gray, 1872a is an unused mis". Spicules are reported to be oxeas of 4250 X 36 p.m, and sig- name in the sense of ICZN Article 23.9 (not used after 1899), it is maspires of 8 p,m. Topsent (1894c) erected this genus in a footnote here suppressed in favour of Tetillidae (used many times by many discussing the genus Gellius (Haplosclerida), assuming apparently authors in the past 50 years). that the microscleres of T globosa were true sigmas. However, it is The genus Chrotella Sollas, 1886a was erected for type clear from Carter's description that they are sigmaspires and that species Chrotella simplex Sollas, 1886a: 181; Sollas, 1888: 17, T. globosa belongs to Tetilla or possibly Craniella, and in view of pi. II figs. 1^1 (by subsequent designation by Sollas, 1888: cxxv), the fact that special cortical oxeas were not reported, it is here con- South Australia, 274 m. The type series (BMNH 1889.1.1.12, with sidered a junior synonym of Tetilla. This assumption has been slides BMNH 1894.11.16.27-32, not examined) consists of four recently verified by re-examination of the type material (slides AM cream-coloured subspherical specimens of up to 2 cm in diameter. G2751-2) by J.N.A. Hooper (courtesy of Dr Penny Berents, AMS), Oscules small, pores in sieve-like areas overlying subdermal confirming its allocation to Tetilla and not to Craniella. spaces, surface uniformly pilose. Sollas records a cortex but neither The genus Ectyonilla Ferrer-Hernandez, 1914a: 452 was his detailed description nor his drawings show a clear cortical spe- erected for type species Tetilla truncata Topsent, 1892a: 36, cialization. There are many subdermal cavities but no peripheral pi. VIII fig. 7 from the NW coast of Spain (by original designation). collagenous region, nor cortical spicules. Megascleres are protri- The genus was assigned to a new family Ectyonillidae defined as aenes, with rhabds of up to 3400 X 20 |xm, cladi up to 158 X 16 p,m; "Sigmatophora with a skeleton of Axinellidae". The holotype is anatriaenes up to 5300 X 16 |xm, cladi up to 47 X 12 |xm; oxeas up housed in MOM (not examined). According to Topsent's descrip- to 3000 X 24 p.m. Microscleres sigmaspires 12 p.m. The characters tion, this is a fragment of a definite Tetilla, with a radiate skeleton of C. simplex appear to overlap with those of Tetilla and accord- of styles of 1200p,m instead of oxeas (probably oxea-derived), and ingly it is considered a junior synonym. anatriaenes and protriaenes. No microscleres. The styles are The genus Spiretta Lendenfeld, 1888: 42 was erected for type unusual, but have nothing in common with those of Axinellidae. species Spiretta raphidiophora Lendenfeld, 1888: 43 (subsequent It is clear that it is a commonplace tetillid, and we assign it to the designation herein, lectotype in AM G9076, designation herein, with synonymy of Tetilla on account of the absence of special cortical a small fragment in BMNH 1886.8.27.634, here designated paralec- megascleres. totype). The description of Lendenfeld makes it likely that this is a The genus Kaira de Laubenfels, 1936a: 175 was erected for species of Tetilla, and Lendenfeld admitted as much himself at a later type species (by original designation) Tethya hebes Lendenfeld, date (Lendenfeld, 1903). Although he mentions a 'cortex' this was 1907: 98, pi. XVI figs 19-38 from NW Australia (type 2MB 768 apparently used to indicate the peripheral skeleton, rather than a real (not examined), with fragment BMNH 1908.9.24.66). This species cortex. The species has an unusual spiculation by its complement of was assigned into synonymy with Cinachyra australiensis (Carter, styles of 600 X 70 p,m, small choanosomal oxeas of 240 X 20 |xm, 1886b: 127 as Tethya cranium var. australiensis) by Burton and ectosomal strongyles of 400 X 3 p,m. Furthermore there are radi- (1934a: 524), followed by Hooper & Wiedenmayer (1994: 430), ating oxeas 3000-4000 X 50 p,m, anatriaenes 1000 X 10p,m, sig- because both possessed rugose small oxeas. However, several dis- maspires 14p,m; apparently no protriaenes. Spiretta was relegated crepancies between the descriptions of australiensis and hebes are into synonymy with Cinachyra by Burton (1934a: 524), as he apparent (presence of plagiotriaenes and anatriaenes in the latter). assumed that the apertures of the type specimen represented 'col- Moreover, Lendenfeld's specimen possessed only a single lateral lapsed porocalices'. This synonymy assignment was followed by oscule and no other apertures, whereas C. australiensis, which is Hooper & Wiedenmayer (1994: 430). Conversely, we prefer here to an obvious representative of Cinachyrella, has abundant porocal- consider Spiretta raphidiophora a member of the genus Tetilla on ices. Until fresh material has been found, we prefer to consider account of the absence of clear porocalices and lack of cortex. Tethya hebes a member of Tetilla.